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BULLETIN DE L'INSTITUT ROYAL DES SCIENCES NATURELLES DE BELGIQUE, BIOLOGIE, 59: 163-168, 1989 BULLETIN VAN HET KONINKLIJK BELGISCH INSTITUUT VOOR NATUURWETENSCHAPPEN, BIOLOGIE, 59: 163-1 68, 1989

A new of the rufipes-complex (Reptilia ), from Northern Madagascar

by Mathias LANG & Wolfgang BOHME

Abstract BoETTGER's Catalogue), a female specimen as lectotype of Zonosaurus rufipes. This specimen, however, by no The taxonomic status of Zonosaurus rufipes and var. subunicolor are means matches BOETTGER's original type description reviewed. In addition, a new species of Zonosaurus closely related to as far as measurements are concerned. First of all the rufipes is described from northern Madagascar. Phylogenetic affinities of this new species are discussed. This study also represents the first lectotype has a SVL of85 mm (total length 205 mm) in report of mite pockets in Gerrhosauridae. contrast to the described SVL of 55 mm with a total Key-words: Gerrhosauridae; Zonosaurus brygooisp. nov.; phylogene­ length of 162 mm (see also BRYGOO, 1985). Rather tic affinities. SMF 40747 of BOETTGER's (188la) original syntype series better fits the measurements given in the type Resume description. The latter specimen also resembles the Le statut taxonomique de Zonosaurus rufipes et var. subunicolor est illustration by BOETTGER (reprocuded in ANGEL, examine. U ne nouvelle espece, qui est proche de Z. rufipes est decrit qui 1942). The designated lectotype has very faint lateral se trouve sur les lles deN osy Beet Nosy Boraha eta Sakana. La position throat stripes, the medial ones entirely absent in systematique de cette nouvelle espece est discutee. Cette etude rap porte contrast to the well-defined 6 throat stripes so charac­ aussi pour Ia premiere fois sur des poches d'acariens dans Ia famille Gerrhosauridae. teristic of rufipes. MERTENS' (1967) lectotype designa­ Mots-clefs: Gerrhosauridae ; Zonosaurus brygooi sp. nov.; implications tion is maintained, bearing in mind that the lectotype is phylogenetiques. considerably larger than the dimensions given in the original type description. Introduction In a recent review of Zonosaurus, BRYGOO (1985) supported the validity of rufipes, rejecting earlier The island of Nosy Be (293 km2) has been a well­ statements by MOCQUARD (1909: 25), who considered known and favorite collecting area since the late 19th rufipes identical to aeneus. ANGEL (1942: 95) on the century(BOETTGER, 1880, 1881a, 1881b). Nevertheless, other hand, doubted the validity of aeneus, but only recently three new saurian species have been presented a list of characters separating rufipes from discovered occurring on this small island belonging both aeneus and madagascariensis. Here we accept respectively to Brookesia(RAMANANTSOA, 1979), Phel­ BRYGOO's (1985) views, because he clearly demonstra­ suma (MEIER, 1988) and Uroplatus (B6HME, in prep.). ted that aeneus, rufipes and madagascariensis constitute It is furthermore surprising that on this island no less valid species. than six species of Zonosaurus occur: aeneus, boettgeri, The taxonomic history of Zonosaurus rufipes var. laticaudatus, madagascariensis, rufipes (see BRYGOO, subunicolor is rather complex. BOETTGER (1881a) 1985 ; MEIER 1989) and the new species described based the description of his var. subunicolor on two below. specimens. The specimen for which accurate measure­ BOETTGER (188la) described the seemingly endemic ments are given (SMF 41051 ; lectotype) as well as the Zonosaurus rufipes with diagnostic throat stripes, and paralectotype (SMF 41052) are clearly Zonosaurus var. subunicolor without throat pigmentation. Taxono­ rufipes based on morphometric characters (Table 1). mic confusion followed the original type description Both have only very faint lateral throat stripes, hence and we therefore wish to reiterate some of the problems the name subunicolor. and clarify the situation. BOETTGER (1913) indicated that the color pattern of Taxonomic account subunicolor may be the juvenile pattern of rufipes. MERTENS ( 1967) rejected this notion and after designa­ MERTENS (1967) designated SMF 40743 (6128, 1a of tion of lectotypes for these two taxa (see above) placed II

164 M. LANG & W. BOHME

Table 1 :Summary ofmeristic values differentiating Zonosaurus on Table 2: Meristic valus ofZonosaurus rifipes. For abbreviations see Nosy Be. For more detail on values ofZonosaurus rufipes and Z. Table 1. brygooi see Table 2 and 3 respectively. A slash mark(/) distinguishes between left and right sides whereas a hash mark (-) indicates a TAXON FEM.POR. SO IP SVL(T01) MID BOD CH-CL 4 TOE range. AMNH 24769 ll fl2 3 PR 35 ('!!) 24 46 21 BM 86.2.25.8 11 /12 3 AB 75 (?!!) 25 48 21 TAXON FEM.POR. SO IP SVL(T01) MlDBOD CH-CL 4TOE ** BM 86.2.25.9 ** 11 / 11 3 AB 51(?!!) 24 48 21 BM 87.12.5.13 11 / 11 3 PR 37 ('!!) 25 49 20 BM 95.10.29.12 11 / 11 3 AB 66('!1) 24 45 20 aeneus CAS 156896 10/ 11 3 PR 31(90) 25 47 21 BRYGOO (1985) 12-19(15) AB 86(236?) 19-23(20) 42-58(51) 16-22( 19) CAS 156897 14/14 3 PR 38 (?!!) 27 48 21 NMW 12243 12/ 12 3 PR 55 (135) . 24 50 22 boettgeri NMW 12245 .1 12/12 2 PR '!! (?!!) 26 46 22 NMW23348 16/ 17 4 PR 113 14 48 22 NMW 12245.2 11 / 11 3 PR 52(?!!) 26 46 22 holotype NMW 12246. 1 13/ 11 3 PR 81 (?!!) 24 '!! 22 NMW 12246.2 12/ 11 3 PR 40 (?!!) 25 48 21 madagascariensis NMW 12247.1 11 / 11 3 AB 69 (197) 25 47 21 BRYGOO (1985) 15-23(19) 4 AB 160 18-24{22) 52~ 57) 20-24{22) NMW 12247.2 9/9 3 AB 69 (195) 26 '!! 21 NMW20097.1 10/ 11 3 AB 80 (204) 25 48 21 laticaudatliS NMW 20097.2 13/14 3 PR 73 (207) 25 51 23 BRYGOO (1985) 17-28(23) 4 AB 142(434) 22-26(24) 45-50(48) 22-27(24) NMW 23350.1 ** 11 / 10 3 PR 34 (?!!) 25 48 22 NMW 23350.2 ** 11 / 11 3 PR 51 (126) 25 48 22 rufipes SMF 40743 * 13fl3 3 AB 85 (?!!) 25 '!! 22 TABLE2 9-14{11) 3 PR 31-85 23-28(25) 44-51(47) 20-23(21) SMF 40744 ** 11 / 10 3 PR 81 (?!!) 27 48 21 SMF 40745 ** 11 / 11 3 PR 44 (99) 25 45 22 brygooi SMF 40746 ** 11 / 10 3 PR 37 ('!!) 24 48 21 TABLE 3 16-19(17) 3 AB 51-76 20-23(21) 43-48(45) 17-19(18) SMF 40747 ** 9/9 3 PR 55(??) 23 46 21 SMF 40748 ** 10/ 10 3 PR 42 (??) 24 44 20 SMF 4105 1 jj 10/ 10 3 PR 45 (142) '!! 46 22 (Abbreviations used inT able 1-4& Fig. I) : Fern. Por. =number of femoral pores on thighs; SMF 41052 !Ill 11 / 11 3 PR 38 ('!!) 25 45 20 SO= the number of supralabial scales anterior to the largesupraocu1ar scale that forms part ZFMK 21270 11 /12 3/4 PR 45 (10 1) 28 50 22 of the labial margin (not including rostral) ; IP =interparietal scale (AB = absent ; PR = ZFMK 46796 10/ 11 3 PR 36 (106) 24 '!! 22 present); SVL =snout vent length, TOT= total length (expressed in mm) ;? =tail incomplete ZFMK 46797 11 /10 3 PR 34 (?!!) 25 47 22 or missing; MID BOD= number of scales at midbodyexcluding the ventral scales ; Ch-Cl = ZFMK 7295 11 / 11 3 PR 36 (102) 25 49 22 nu mber of scales between chin and cloaca (including mental scale) ; 4 toe = number of ZFMK 48239 13/12 3 PR 36 (100) 27 49 22 subdigital scales below 4th toe. ZMB 10097 ** 14/14 3 PR 63 (109) '!! 47 21 ZMB 10097 ** II JI2 3 PR 48 (76) 25 47 21 RANGE: 9-14 31-85 23-28 44-51 20-23

MEAN: 11.3 25.1 47.4 21.5 var. subunicolor as a synonym of rufipes. Finally, * = lectotype Zonosaurus rufipes BRYGOO, 1985:34) categorically states that from a ** = paralectotype ZonosaurliS rufipes taxonomic standpoint subunicolor can not be distin- !I = lectotype Z. rufipes var. subunicolor guished from rufipes and is therefore considered to be a !Ill = paralectotype Z. rufipes var. subunico/or junior synonym thereof. After examination of new material from Nosy Be in addition to the type material of Zonosaurus rufipes and Table 3: Meristic values ofZonosaurus brygooi. For abbreviatons see Table 1. subunicolor, it is evident that the description of a new species of Zonosaurus is warranted. TAXON FEM.POR. SO lP SVL(T01) MID BOD CH-CL 4 TOE ZFMK 46789 * 19/19 AB 76 (?!!) 22 44 18 Zonosaurus brygooi sp. nov. ZFMK 46790 ** 17/ 17 AB 63 (?!!) 21 43 19 ZFMK 46792 ** 16/16 AB 51 (145) 23 45 17 Diagnosis. - Zonosaurus brygooi differs from other ZFMK 46793 ** 19/ 17 AB 55(?!!) '!! '!! 18 species of Zonosaurus by the absence of an interparietal ZFMK 46794 ** 17 / 16 AB 45 (129) 21 46 18 ZFMK 46795 scale, relatively large dorsal scales (20-23), few scales ** 16/ 16 AB 49 (124) 20 48 18 ZFMK 48165 ** 16/ 16 AB 74 (?!!) 22 45 18 between chin and cloaca (43-48), a relatively high IRSNB 2.534 ** 17 f1 7 AB 74 (?!!) 22 45 18 number of femoral pores (16-19) and few subdigital ZMB 190 18 ** 16/17 AB 69 (161) 21 45 17 lamellae below the 4th toe (17-19) (Fig. 1-4; Table 1). SMF 41053 ** 17/1 7 AB 68 (?!!) 21 47 19 Zonosaurus brygooi distinguishes itself from Z. rufipes RANGE : 16-19 51-76 20-23 43-48 17-1 9 by the consistent lack of throat striping, interparietal scale and 3 characteristics of lepidosis that do not MEAN: 16.9 21.4 45.3 18 overlap (Fig. 1-4; Table 1). Zonosaurus rufipes and * = holotype brygooi can furthermore be distinguished from the ** = paratype remaining Zonosaurus (except quadrilineatus and trili- neatus) by having an entirely dark pigmented tongue. 'I

A new species of Zonosaurus 165

FEM. PORES SCALES AT MIDBODY I • • II • II • Ill • Ill • IV • IV 2 8 • -- -j v v • VI VI •

9 11 13 15 17 19 21 23 14 16 18 20 22 2 4 26 2 8 Fig. 1.: Diagram illustrating differences in the number offemora l pores between the 6 species ofZonosaurus occurring on Nosy Be. /) brygooi; II) rufipes;2 III) aeneus; IV) madagascariensis; V) laticaudatus and VI) boettgeri. (Values derived from Table 1).

Fig. 2.: Diagram illustrating differences in the number ofsca les at midbody between the 6 species ofZonosaurus occurring on Nosy Be. I) brygooi; II) rufipes; Ill) aeneus; IV) madagascariensis; V) laticaudatus and VI) boettgeri. (Values derived from Table I).

SUB DIG. LAM. 4TH TOE SCALES CHIN - CLOACA • • II II • • 16 Ill ~-- - Ill IV IV • • 2 7 v • v • --1 VI • VI •

43 17 18 19 2 0 21 22 23 2 4 Fig. 3.: Diagram illustrating differences in the number ofscales between the chin and the cloaca ofthe 6 species ofZonosaurus occurring on Nosy Be. I) brygooi; II) rufipes; Ill) aeneus; IV) madagascariensis; V) laticaudatus and VI) boettgeri. (Values derived from Table 1).

Fig. 4.: Diagram illustrating differences in the number ofsubdigitallamellae below the 4th toe in the 6 species ofZonosaurus occurring on Nosy Be. I) brygooi; !I) rufipes; II/) aeneus; IV) madagascariensis; V) laticaudatus and VI) boettgeri. (Values derived f rom Table 1).

Zonosaurus aeneus, brygooi and rufipes have 3 supra­ Description ofholotype. - (Fig. 5-6). Nasal scales not in labial scales anterior to the sub ocular that forms part of median contact; separated by large frontonasal scale. the upper labial margin, whereas the remaining Zono­ Prefrontals separated by frontal-frontonasal contact. saw·us all have 4 supralabials anterior to the subocular. Two large parietal scales; no frontoparietals. Interparie­ In addition, these three species all have 2-3 mite pockets tal scales absent. Four supraocular scales. Four supraci­ located in the antehumeral fold in addition to postaxil­ liary scales. Two loreals. A total of 6 scales between lary and postfemoral mite pockets. This condition is rostral and tympanum with 3 scales anterior to the not found in any other species of Zonosaurus. The mite subocular, the 4th scale being the subocular. A total of pockets contained larvae of Ophionyssus natricis (Lae­ 5 temporal scales. Lower eyelid contains about 13 laptidae). For differentiation from the remaining Zono­ scales forming a translucent window. Temporal and saw·us see BRYGOO (1985). parietal scales with slight carinations (Fig. 5). Lateral and dorsal body scales have slight carinations. Holotype. - ZFMK 46789, a male from Loucoube, The number of midbody scales, scales between chin Nosy-Be, Madagascar; coli. R. Seipp, IV 1987. and cloaca, subdigital scales below the 4th toe as well as \I

166 M. LANG & W. BOHME

lighter bluish-white flecks of the longitudinal lines are lateral to the darker flecks. The longitudinal lines stop short of the sacral region. Also on the flanks are up to 13 light blue spots. A few of these spots are located on the dorsal aspect of the anterior limbs. Darker pigmented areas are found in the dorsum, and in the neck region, where up to 6 spots form a short interrupted line. These spots also continue into the tail region.

Paratypes. - ZFMK 46790; 46792-46795 same data as holotype; IRSNB 2.543 same data as holotype; ZFMK 48165 Madagascar, Nosy Boraha (== Ile Ste. Marie), coli. F.W. Henkel & R. Seipp, IV 1988; ZMB 19018, Madagascar, Sakana, coil. Voeltzkow, no date; SMF 41053 Nosy Be, Madagascar, coli. A Stumpff, 1881. The latter specimen was listed as Zonosaurus aeneus by BRYGOO (1985: 44).

Variation. - Paratypes for the most part follow the description of the holotype. For variation of lepidosis see Table 3. No considerable color variations of paratypes with respect to the holotype were observed. MEIER ( 1989: Fig. 8 a, b) presents color photographs of the underside of both Z. brygooi and Z. rufipes, showing the striking color and pattern differences between the two species. ZFMK 46790 is remarkable in Fig. 5.: Upper and lateral views ofthe head ofthe holotype (ZFM K 46789) ofZonosaurus brygooi sp. nov. that it.has the head of a swallowed juvenile protruding from its mouth. This was terrarium kept and it the number of femoral pores are listed in Table 3. At is therefore not known if cannibalism is a natural midbody the ventrals are arranged in 8 longitudinal behavior of this species. rows. The tail is abruptly constricted just posterior to the sacral region, whereafter it is vertically compressed Distribution and Habitat. - Zonosaurus brygooi occurs (Fig. 6). The scales on the distal aspect of the tail are on the island of Nosy Be and Nosy Boraha and at smooth. Sakana (Eastern Forest). The latter locality is proble­ matic and has not been found on any map or in any Head : II mm; Trunk: 65 mm; Snout-vent length: 76 gazetteer (BRYGOO and BOUR, pers. comm.). We mm; head width at tympanum: II mm; anterior limb: suspect that it is located on the north eastern coast of 23 mm; posterior limb: 41 mm; incomplete tail: 66 mm. Madagascar, coinciding with the travel routes of The holotype is a well-preserved male specimen with VOELTZKOW. Sakana was cited by BRYGOO (1985) as non-faded colors and a snout-vent length of 76 mm. being erroneous for Zonosaurus nifipes. Indeed the The head is a metallic medium brown with some specimen from that site is Z. brygooi. This new species iridescence. A few dark pigmented areas are situated occurs in regions of primarily damp wet rainforests along the lateral aspect of the supraoculars and below (MEIER, 1989; listed as Zonosaurus rufipes). the eye. The supralabials are only slightly pigmented. As can be deduced from the geographical distribution The throat and chest areas are a light blue, which of Zonosaurus, some species show a disjunct distribu­ grades to a light red in the abdominal area, cloacal tional pattern between the forest relicts of Nosy Be and region and interior thighs. The palmar surface is also the forested areas of northeastern Madagascar, inclu­ reddish (as is the case in Zonosaurus rufipes). The ding the island of Nosy Boraha (= Ile Ste. Marie). ventral aspect of the tail is light blue. Species exhibiting this pattern include aeneus, boettgeri The lateral aspect of the body, the dorsum and the and brygooi. Zonosaurus rufipes is also a forest dorsal aspect of the tail are a darker shiny brown. The inhabitant and has been found at Imerina (Eastern flanks are separated from the dorsum by 2 interrupted Madagascar). Zonosaurus laticaudatus and madagas­ longitudinal lines that start just posterior to the parietal cariensis, which are adapted to more open habitats region. These lines consist of bluish-white and deep show a more continuous distribution throughout the brown rectangles I scale wide and 2 scales long. These main island of Madagascar (BRYGOO, 1985: 26 & 56). longitudinal lines are interrupted by scales (usually I) The niche segregation and resource partitioning of having the dorsal medium brown coloration. The these six species of Zonosaurus on the small island of '' A new species of Zonosaurus 167

.,

' ! I /I ., i I' '"1""1""'" II III II I' II 'I" II jiiiiJ II II IIIIIJIIIIjiiiiJIIIIjllllllllllllll!ll lllllll !I I i I i I" I I ' : 7 8 9 . 0 1 2 3 4 5 6 7 8

Fig. 6.: Dorsal and venlral views of the holotype ofZonosaurus brygooi.

Nosy Be would constitute a most interesting ecological phylogenetic results indicate that aeneus + (rufipes + project. brygoo1) form a phylogenetic unit that is based on the presence of 2-3 well-defined mite pockets within the Phylogenetic affinities. - Zonosaurus brygooi is more antehumeral fold and in having 3 supralabial scales closely related to Z. rujipes than to any other species of anter~or to the subocular scale that is part of the labial Zonosaurus. This is based on the presence of an entirely margm. pigmented tongue (convergent with Z. trilineatus and Z. quadrilineatus) and the possession of reddish feet, Etymology. - This new species is named in honour of which is not found in any other species of Zonosaurus. Professeur E. R. BRYGOO who has contributed so The entire is at present being analyzed within a much to the knowledge of the fauna of Madagas­ phylogenetic framework (LAN G, in prep.). Preliminary car, in particular of the genus Zonosaurus. I I

168 M. LANG & W. BOHME

Acknowledgments. drew STIMSON & Colin McCARTHY (BM), Richard ZWEIFEL (AMNH) and Jose ROSADO (MCZ). Dr. We wish to thank the following persons for the loan of Alex FAIN is thanked for the identification of the material under their care: Jens VINDUM (CAS), Ramer trombiculid larvae. Many thanks also to an anony­ GUNTHER (ZMB), Konrad KLEMMER and Monika mous reviewer for constructive criticism. LAUDAHN (SMF), Franz TIEDEMANN (NMW), An-

Literature cited

ANGEL, F., 1942. Les Lezards de Madagascar, Memoires de MEIER, H., 1989. Zur Okologie, Ethologie und Taxonomie L'Academie Malagache, 36: 1-194. einiger Schildechsen der Gattung Trachelop tychus und Zonosaurus auf Madagaskar. Teil 2: weitere Zonosaurus BdHME, W., (in prep.). Urop/atusflmbriatus complex. -Arten. He1petojauna, 11(58): 14-23.

BOETIGER, 0., 1880. Diagnoses reptilium et batrachiorum MERTENS, R., 1967. Die herpetologische Sektion des Natur­ novorum a Carole Ebenau in insula Nossi-Be Madagas­ Museums und Forschungsinstitutes Senckenberg in Frank­ cariensi lectorum. Zoologischer Anzeiger, 57: 279-283. furt a. M . nebst einem Verzeichnis ihrer Typen. Sencken­ bergiana Biologica, 48A: 1-106. BOETIGER, 0., 1881 a. Diagnoses reptilium et batrachiorum MOCQUARD, F., 1909. Synopsis des families, genres et novo rum ab ill. Antonio Stumpff in insula N ossi-Be Mada­ especes des ecailleux et des Batraciens de Madagas­ gascariensi lectorum. Zoologischer Anzeiger, 4: 358-362. car. Nouvelles Archives du Museum National d 'Histoire Natt.irelle, Paris, 5e S(l ): 1-108. BOETIGER, 0., 1881b. Die Reptilien und Amphibien von Madagascar. Dritter Nachtrag. Abhandlungen der Sencken­ RAM ANANTSOA, G., 1979. Description de deux nouvelles bergischen Naturforschenden Gesellschaft, 12: 435-558. especes de Brookesia (Reptilia, Squamata, Chamaeleonidae). B. legendrei et B. bonsi. Bulletin du Museum National BOETIGER, 0 ., 1913. Reptilien und Amphibien von Mada­ d'Histoire Nature lie, Paris, 4e ser., 1, 1979, section A, 3: gascar, den Inseln und dem Festland Ostafrikas. In: Reise in 385-693. Ostafrika in den Jahren 1902-1905 ... Alfred Voeltzkow 3(4).

BRYGOO, E.R., 1985. Les Gerrhosauridae de Madagascar. Mathias LANG* and Sauria (). Memoires du Museum National d'His­ Wolfgang BOHME toire Naturelle, Paris. SerA, Tome 134: 1-65. Herpetologische Abteilung Zoologisches Forschungsinstitut LEVITON, A. E., McDIARMID, R., MOODY, S., NICKERSON, und Museum Alexander Koenig M., ROSADO, J., SOKOL, 0. & VORIS, H. 1980. Museum Adenauerallee 150-164 acronyms - second edition. Herpetological Review, 11(4): D-300 Bonn 1, West Germany 93-102. *Research Associate Koninklijk Belgisch Instituut MEIER, H., 1988. Vorlaufige Beschreibung einer neuen Art voor Natuurwetenschappen der Gattung Phelsuma von Madagaskar. (Sauria: Gekkoni­ Vautierstraat 29 dae). Salamandra, 23(4): 204-211. B-1040 Brussels, Belgium

Specimens examined 12245.1-2, 12246.1 -2, 12247. 1-2, 20097. 1-2, 23350. 1-2 (para­ lectotypes) Nosy Be; SMF 40743 (lectotype), 40744-40748 (para­ For museum acronyms see LEVITON et al. (1985). lectotypes), 41051 (lectotype of var. subunicolor), 41052 (paralecto­ Zonosaurus brygooi: type) Nosy Be; ZFMK 21270 no data; ZFMK46796-46797,47295, Holotype: ZFMK 46789, Nosy Be, Loucoube 48239 Nosy Be, Loucoube; ZMB 10097 (2 spec.) (paralectotypes) Paratypes: ZFMK 46790, 46792-95, Nosy Be, Loucoube; SMF Nosy Be. 41053 Nosy Be, IRSNB 2.534 Nosy Be, Loucoube; ZFMK 48165, Zonosaurus boettgeri: NMW 23348, Nosy Be, 189 1. Nosy Boraha (= Ile Ste. Marie); ZMB 1901 8, Sakana. Zonosaurus aeneus: ZFMK 14365, 21272, Nosy Be (Madagascar). Zonosaurus rufipes: AMNH 24769 Nosy Be; BM 86.2.25.8-9 Nosy Be (paralectotypes); BM 87. 12.5. 13 Nosy Be; BM 95. 10.29. 12, Remaining Zonosaurus examined are from the collections of lmerina, Madagascar. CAS 156896-156897 Nosy Be, Lokobe ZFMK (Bonn), IRSNB (Brussels), SNMS (Stuttgart), SMF Forest; MCZ 17633 Nosy Be (paralectotype); NMW 12243, (Frankfurt) and ZMB (E. Berlin).